Dissertation Proposal - The University of Arizona Campus Repository
Dissertation Proposal - The University of Arizona Campus Repository
Dissertation Proposal - The University of Arizona Campus Repository
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I - INTRODUCTION<br />
<strong>The</strong> fitness <strong>of</strong> any evolutionary unit can be understood in terms <strong>of</strong> its two basic<br />
components: fecundity and viability. As embodied in current theory, the trade-<strong>of</strong>fs<br />
between these fitness components drive the evolution <strong>of</strong> life-history traits (Stearns 1992).<br />
In unicellular individuals, the same cell must be involved in both fitness components,<br />
typically these components being separated in time. However, in multicellular organisms,<br />
under certain circumstances, cells may specialize in one component or the other, the<br />
result being a division <strong>of</strong> labor, leading to the differentiation <strong>of</strong> germ and soma. <strong>The</strong><br />
evolution <strong>of</strong> a specialized and sterile soma can increase viability and indirectly benefit<br />
fecundity (e.g., increasing nutrient uptake) but, all things being equal, must directly cost<br />
fecundity by reducing the number <strong>of</strong> cells producing <strong>of</strong>fspring. On the other hand, the<br />
evolution <strong>of</strong> a specialized germ will benefit fecundity (by reducing the generation time<br />
and/or increasing the quality <strong>of</strong> <strong>of</strong>fspring), but must directly cost viability by reducing the<br />
number <strong>of</strong> cells participating in viability-related functions.<br />
Various selective pressures may push unicellular organisms to increase in size, but<br />
general constraints, such as the decrease in the surface to volume ratio, set an upper limit<br />
on cell size. Increase in size can also be achieved by the aggregation <strong>of</strong> mitotic products<br />
that are held together by a cohesive extra-cellular material, increasing the number <strong>of</strong> cells<br />
(instead <strong>of</strong> cell size). Natural selection has favored this strategy as illustrated by the<br />
multiple independent origins <strong>of</strong> colonial and multicellular organisms in, for example,<br />
algae (Niklas 1994; 2000). Large size can be beneficial both for viability (e.g. in terms <strong>of</strong><br />
predation avoidance, ability to catch bigger prey, a buffered environment within a group)<br />
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